February 25, 2014

Recent radiation of R-M269 males in Europe

From the paper:

As such it is not possible to predict the subhaplogroup within R-M269 to which an individual belongs based on his YSTR haplotype, in contrast to the situation with higher level haplogroups for which haplotypes do have predictive power(Athey, 2005; Schlecht et al., 2008).

The explosion of R-M269 descendants who (seemingly) went from zero to a solid majority in much of Western Europe over the last few thousand years is probably one of the most interesting events in recent European history. Not many would have entertained such a possibility until a few years ago, but truth is often stranger than fiction.

Y-chromosomal short tandem repeats (Y-STRs) are often used in addition to Y-chromosomal single-nucleotide polymorphisms (Y-SNP) to detect subtle patterns in a population genetic structure. There are, however, indications for Y-STR haplotype resemblance across different subhaplogroups within haplogroup R1b1b2 (R-M269) which may lead to erosion in the observation of the population genetic pattern. Hence the question arises whether Y-STR haplotypes are still informative beyond high-resolution Y-SNP genotyping for population genetic studies. To address this question, we genotyped the Y chromosomes of more than 1000 males originating from the West-European regions of Flanders (Belgium), North-Brabant and Limburg (the Netherlands) at the highest resolution of the current Y-SNP tree together with 38 commonly used Y-STRs. We observed high resemblance of Y-STR haplotypes between males belonging to different subhaplogroups of haplogroup R-M269. Several subhaplogroups within R-M269 could not be distinguished from each other based on differences in Y-STR haplotype variation. The most likely hypothesis to explain this similarity of Y-STR haplotypes within the population of R-M269 members is a recent radiation where various subhaplogroups originated within a relatively short time period. We conclude that high-resolution Y-SNP typing rather than Y-STR typing might be more useful to study population genetic patterns in (Western) Europe.

53 comments:

So, if I understand this correctly, men of diverging subhaplogroup lineages have a good chance of undergoing the same (or very similar) STR mutations. In other words, those are not random, but occur over and over again, at least for members of a specific haplogroup (here, M269). One would expect that the farther removed men are in terms of their haplogroup, the more random the STR variations - which then makes them more useful to distinguish haplogroups at the higher level.

I guess there is something to be said for checking and publishing a peer reviewed confirmation of something which was widely understood by people working practically. OTOH, I guess the obvious rejoinder is that you could use more STR markers, and test again. Genealogist admins have been telling R-M269 people for many years to get as many as they can afford and 111 is quite a standard test these days.

Still doesn't really add up. Repeats move both ways, and we don't know the range is unconstrained, so this kind of logic simply can't be used to come to a solid conclusion.

The math has to make assumptions we can't make, therefore we just can't use that math for this purpose.

The shear number of subtypes or r1b also says that expansion was not nearly so rapid. If they truly shifted all around at random, then they would be distributed in a uniform pattern. Clearly these are separate clades that had to have formed thousands of years ago themselves or have some WILDLY large selection advantage to have geographic based distribution.

This is a case where the apriori probability of STR permutations generating the same results is greater for different R1 haploid groups than others. Before recent extensive SNP Y dna testing the 111 STRs, not 38, were useful in differentiating some haploid groups in I for which there were no known differentiating SNPs. These other haploid groups have different a-priori probabilities of having the same STR permutations.

This is why popular y-dna estimates of genealogical relationships based on STRs are dumbed down for the R1 European majority, while for other haploid groups people in the same family tree are related with as many as 7 differences in 37 STRs.

Brace for a marketing program to sell SNP instead of STR testing to the R1 majority.

Andrew, R1b in west Europe did not expand in the last 2,000 years. It was already standard by the time of the Roman empire(and had been for hundreds or even a 1,000 years). Since the last 2,000 years is historical times, who spread these young branches? Obviously there is no logical answer and all or most of the expansion was pre historic.

i'm with phil said:"Over the last few thousand years? More like 5,000-3,000BP so for the last 3,000 years for most of west Europe. You make it seem like R1b L11 expanded during the Roman empire and middle ages."

and Grognard: "The shear number of subtypes or r1b also says that expansion was not nearly so rapid. If they truly shifted all around at random, then they would be distributed in a uniform pattern. Clearly these are separate clades that had to have formed thousands of years ago themselves or have some WILDLY large selection advantage to have geographic based distribution."

Since I was around 10 they went from 80,000 all the way down to 13,000 years for native americans. I guess they actually started at 130,000 years. Now it's 4,000 for the east coast as some claim to have found european dna on the east coast that predates natives there...We've always felt it was a stretch to accommodate the genocides that occurred in the americas.

So if they stretched it from 80k or 130k before that all the way down to 13k or 4k then I would also expect them to attempt to embellish on the other end as well, that is raising the age on the other end as much as possible. So I wonder if there's a tendency to look for or "make" evidence to do that. Also I think allot more people in the USA and Europe have taken the tests so that also may effect "the shear number of subtypes".

barakobama, with 67 markers (which is more than were used in the paper) you have a very good chance of being able to get a GOOD prediction of your SNP status, if you consult with people familiar with the typical STR haplotypes of the SNPs you are interested in. Presumably there is a haplogroup project you can contact, and they should be able to help you. This article should not be read as saying otherwise in my opinion.

Do they really say that the mutations of different lineages are non-random and likely to be similar in related lineages?

Andrew - no, that is my interpretation. They claim the possibility of identity by descent. But that is IMO not possible, at least not this extreme. It would require that several to many thousand years ago, there was a fairly large M269* population that had acquired hundreds of different STR mutations, but no surviving SNP, yet*. Then it expanded and acquired dozens of SNP mutations - but with relatively few STR changes. That is impossible given the much higher STR mutation rate. It would only be possible if a good fraction of the STR mutations went back and forth very similar paths in the network in the figure Dienekes provides. But if that is the case, the simpler explanation is that this happened throughout, from the beginning. In other words, that the paths in the network are non-random for a specific, narrow haplogroup.

For example, there's lots of black z381 dots all around the edges of the network, in conjunction with not just the relatively closely related pink U198 cluster and orange U106 but also the remote gray U152. Or orange U106 and green P312 who are far apart, etc.

Or, look at the bottom of the network where there is a transition from (relation between) yellow L48, green P312, then again yellow L48, and finally grey U152 - which does not at all correspond to the SNP tree (Fig. 1).

While M269* may have harbored many different STRs, each color dot must have originated from the same junction of the network. That's easy to see for yellow - but much less so for e.g., green or black without running into contradiction with the SNP tree.

* I would agree that the inner, spidery network of the graph could correspond to that scenario.

We have a number of Mesolithic Y-DNA samples, in which Y-DNA I seems to be predominant. See Lazaridis, I. et al. (2013), Ancient human genomes suggest three ancestral populations for Europeans, pre-print online 23 December 2013. http://biorxiv.org/content/early/2013/12/23/001552 (looking at one sample from European hunter-gatherers Loschbour, Heffingen [LSB 1] in Luxembourg (6220-5990 BCE) and four more European hunter-gatherer samples from Motola, Sweden ca. 6000 BCE.

We have a number of first wave Neolithic Y-DNA samples (both LBK ca. 5500 BCE to 4900 BCE, and the roughly contemporaneous parallel Cardial Pottery Neolithic along Europe's Southern Coast) in which, for example, Y-DNA G is common and R1b is absent.

The earliest ancient DNA evidence of R-M269 (aka R1b1b2) in Europe is found in two Bell Beaker culture men from ca. 2615 BCE +/- about 65 years buried in Kromsdorf, Germany. R1b antecedents to the R1b clade of R-M269 which is predominant in Europe are pretty much absent from Western Europe (both today and in ancient Y-DNA). So, this highly specific subclade of R1b arrived fully developed (this R-M269 clade is, for example, profoundly more distantly related to R-V88 found mostly in Chadic populations in the African Sahel).

The predominance of R1b-M269 in modern Basque populations, the only Western European population not to be converted to Indo-European languages strong suggests that R-M269 predates the arrival of Indo-Europeans in Western Europe.

We have lots of Y-DNA samples from the historic era and the European mix of Y-DNA is pretty stable by not later than 800 BCE or so, maybe earlier. The modern majority view (although certainly not a consensus view) is the Bell Beaker people spoke a non-Indo-European language (probably in the same linguistic family as Basque or proto-Basque), and that Indo-European migration to Western Europe began with the Urnfield culture's arrival around 1300 BCE.

Thus, R1b expansion had probably pretty much run its course by 1300 BCE, there was R1b in Europe in Bell Beaker people by 2650 BCE, and R1b was vanishingly rare in Europe at the tail end of the first wave Neolithic as of 4900 BCE.

There is about a 2250 year window during which the R-M269 expansion could have begun and a 3600 year window between the earliest time it could have begun its expansion and the time when it must have completed it. The fifteen hundred year period from 4800 BP (early Bell Beaker) to 3300 BP (arrive of Indo-Europeans is the most plausible time for that expansion and consistent with this new study. One can't really rule out R1b's expansion with Megalithic cultures ca. 6000 BP to 4800 BP, but these cultures as so deeply intertwined with the first wave Neolithic in Europe, which was followed in many areas by population collapse (sometimes with reversion of foraging) after which there would be a window for resettlement by genetically distinct populations, that this seems less plausible to me.

"The shear number of subtypes or r1b also says that expansion was not nearly so rapid. If they truly shifted all around at random, then they would be distributed in a uniform pattern. Clearly these are separate clades that had to have formed thousands of years ago themselves or have some WILDLY large selection advantage to have geographic based distribution."

There may be a socio-biological explanation for the such a "WILDLY large selection advantage". From the Bronze Age onwards, European societies came to be dominated by chiefdoms based on wealth in cattle and horses. The increased practice of polygamy allowed rich men to not only acquire more wives, but also enabled them to extend their reproductive years into middle age. (For example, the huge number of descendants of Niall of the Nine Hostages) This gave rise to a number of consequences, not the least of which would have been a higher rate of mutations in their offspring, which has been demonstrated a number of studies:

http://www.scientificamerican.com/article/older-fathers-pass-on-mor/

In addition to a higher rate of mutations, the prevalence of polygamy would have created a class of disaffected "bachelor warriors" would would have been highly motivated to attack other groups in order to steal their livestock in order to pay a bride-price, or abduct their women (marriage by capture, as immortalized in Roman history).

An additional consequence of an increased rate of births to older fathers would have been an increase in mental pathology. Studies have shown that children born to older fathers have greatly increased risk of bipolar disorder and depression, as well as ADHD, which, along with the "bachelor warrior" phenomenon, would have driven extremely high levels of inter- and intra-group aggression, leading to expansionist behavior.

@ Andrew Great summary, but"The predominance of R1b-M269 in modern Basque populations, the only Western European population not to be converted to Indo-European languages strong suggests that R-M269 predates the arrival of Indo-Europeans in Western Europe. "

You are presuming Indo-European speakers actually *arrived*. This has been a lasting miscopnception, and I dare say, error propagated by archaeologists, linguists an population geneticists.

We must analyze genetics independently of (hsaky) linguistic hypotheses, and not make circular arguements

'The modern majority view (although certainly not a consensus view) is the Bell Beaker people spoke a non-Indo-European language (probably in the same linguistic family as Basque or proto-Basque)'I am not sure what obsolete Kurganist dwarfs your are referring to. Cunliffe still explicitly left open the possibility that Bell Beaker derives from the westernmost part of the Corded Ware horizon, and most probably Indo-European. This make German-Bell Beaker R1b and German-Corded Ware R1a not only the earliest known carriers of the most prolific modern YDNA R in Europe that are also most commonly associated with Indo European expansion, but also very early Indo European neighbours sharing closely related haplogroups. Moreover, Beaker cultures share a much older common history in this part of the world that has nothing to do with the Neolithic advance whatsoever.

'The predominance of R1b-M269 in modern Basque populations, the only Western European population not to be converted to Indo-European languages strong suggests that R-M269 predates the arrival of Indo-Europeans in Western Europe.'The expansion of YDNA R1b in Europe is indicated as a recent phenomenon. Accordingly, Basque R1b is just a subset of a much wider R1b continuum. All that follows from this pattern of Basque participation in R1b expansion is they must have been on the receiving end.

'R1b was vanishingly rare in Europe at the tail end of the first wave Neolithic as of 4900 BCE.'At least it wasn't found in Mesolithic or LBK Europe up to now. Not any Mesolithic Beaker culture was sampled yet.

'Indo-European migration to Western Europe began with the Urnfield culture's arrival around 1300 BCE.'Hard to disprove on genetic grounds since they burned their dead. But alas, all I can see here is another attempt to dismiss all accumulating evidence, because none of late 20th century Indo European scenarios offer the required flexibiliy and predictive power to cope with modern evidence.

Finally some comments that make sense. Pretty much ditto to everything you said. Thanks for presenting the case and backing up your theories with science.

In sum, all evidence we have currently indicates (notice I said, "indicates") that a small population of R-M269 (or a close predecessor) was somewhere in Western Europe (notice I did not specify Iberia) at the dawn of the Bell Beaker revolution.

It expanded BEFORE the Indo-European invasions.

The discussions of why they were competitively better or something is nonsense. All one needs to know for the expansion is to accept some of the earlier theories that Bell Beaker types were the first who were Lactose Tolerant or the first who drank beer and wine in Europe.

(I mean, think about it. These Bell Beaker people practically worshiped pottery that was likely used for milking or drinking milk. So much so, that they buried their dead with it.)

Read about the Lactose-tolerant mutation and the period from which it derives. This was a time of massive famine in Europe, caused by something akin to a mini-ice age and massive crop failure. There was also a massive decrease in available animal species to be hunted. Population decreased severely, then bottlenecked.

There is your explanation on "how Europe came to be R1b." It was thinly populated to begin with, and then with a famine, some came to be able to digest lactose easily. They owned cattle, they lived.

Several anthropologists have joked that what defines modern Europeans is a smidgen of Neandertal blood, the ability to digest lactose, and a lot of inbreeding. There you have it.

'There is your explanation on "how Europe came to be R1b." It was thinly populated to begin with, and then with a famine, some came to be able to digest lactose easily. They owned cattle, they lived."

Whatever the specifics might be, R1b expansion in WE was all about economy, ecology and demographics.

It had nothing to do with whether they were, or were not, Indo-European speakers.

Any simple equation of 'archaeological cultures' with language relies on an utter ignorance of the complexities of ancient sociolinguistics. Anyhow, the Bell Beaker phenomenon was far too early for PIE in WE. IMHO, PIE in WE was a rather late phenomenon, ie with the advent of proto-Celtic during the Late Bronze/ early Iron Age. !!!

The Beaker culture was also present in North Africa. Sadly we have very few dates for the culture but it is possible that Africans may have taken R-M269 to Iberia/Europe.

Many researchers have committed on the large number of R-V88 who speak Afro-Asiatic languages. But few have noticed that many Niger-Congo speakers, Pgymies and Khoisan carry R-M269.

. Berniell-Lee et al (2009) found in their study that 5.2% carried Rb1*. The frequency of among the Bantu ranged from 2-20. The bearers of R1b1 among the Pygmy populations ranged from 1-25% (Berniell-Lee et al, 2009). The frequency of R1b1 among Guinea-Bissau populations was 12% (Carvalho et al,2010).Gonzalez et al, in The genetic landscape of Equatorial Guinea and the origin and migration routes of the Y chromosome haplogroup R-V88, found that 53& of the subjects in his study carried R-M269.

Given the Corded Ware/Beaker culture sites in North Africa, and widespread evidence of R-M269 from Guinea to South Africa offer the possibility that Sub-Saharan Africans may have introduced this haplogroup to western Eurasia.

"'The predominance of R1b-M269 in modern Basque populations, the only Western European population not to be converted to Indo-European languages strong suggests that R-M269 predates the arrival of Indo-Europeans in Western Europe.'The expansion of YDNA R1b in Europe is indicated as a recent phenomenon. Accordingly, Basque R1b is just a subset of a much wider R1b continuum. All that follows from this pattern of Basque participation in R1b expansion is they must have been on the receiving end."

An Armenian friend of mine recently passed along a link to an article discussing connections between the Basques and the Armenians. The YHG R1b link figured prominently"

For a proximate examination of European affiliations to Armenians, you may also refer to a study done by Michael E. Weale, Rolf F. Jager and Neil Bradman in 2001 called “Armenian Y chromosome haplotypes reveal strong regional structure within a single ethno-national group, revealing prevalent link between Welsh, Basques and Irish to the Armenian populace in Syunik and Karabakh.

The article went on to list several words that were alike in Basque and Armenian, but missed a major point: the word for 'water' in Armenian, jur, closely resembles the Basque equivalent ur, and both appear to be derived from the Yeniseian language family, despite the fact that Armenian is classified as an IE language. Together with hydronymics from the Balkans, the possible footprints of the immigration of the ancient pre-Vasconic peoples from the Caucasus to central Europe might be deduced.

Another interesting feature that potentially links Armenian to both Basque and IE are ending for plural pronouns. For example, IE and Iroquoian languages, the word for 'they' contain the word particle -n- as a marker from plural - for example"

Your mention of the Basque is interesting. This because the Basque language is related to Dogon. Dogon is a Niger-Congo speaking population that lives in Mali, near the Niger River.

It is interesting to note that Africans may have early settled Iberia. Dr. Martin, a tenured professor of Language and Literature at the Instituto Cervantes in Madrid, has spent twelve years to compare Basque and Dogon, both linguistic structure and vocabulary. Professor Jaime Martin, who has compared 2274 words from both languages finding similarities in 70% of them.

"Your mention of the Basque is interesting. This because the Basque language is related to Dogon. Dogon is a Niger-Congo speaking population that lives in Mali, near the Niger River.

It is interesting to note that Africans may have early settled Iberia. Dr. Martin, a tenured professor of Language and Literature at the Instituto Cervantes in Madrid, has spent twelve years to compare Basque and Dogon, both linguistic structure and vocabulary. Professor Jaime Martin, who has compared 2274 words from both languages finding similarities in 70% of them."

I can't confirm that because I have not been able to find a Swadesh list for Dogon language online. Moreover, the affinity with Niger-Congo languages is somewhat tentative. As for genetics, the Dogon exhibit no YHG R1b at all.

My research appears to indicate that much of western Eurasia during the Neolithic and early Bronze Age was dominated by populations which spoke languages related to some modern Native American languages, and which were involved in a sprachbund resulting in extensive linguistic borrowing between groups. This apparent diffusion of peoples out of Siberia and onto the Eurasian steppes presaged the later expansion of the Turks from the Altai region millenia later. The Turks arrived in Anatolia around 1000 AD, but now almost all of the Anatolian population speaks a Turkish language, even though results of genetic testing show they have only a small East Asian component. Likewise, modern Western Europeans speak a language largely derived from a language that was probably also ancestral to Iroquoian, with a contribution from Dene-Yeniseian languages as well, and display only a small residual amount of NA DNA. The parallel I see is with the Garifuna language of the Caribbean. After 500 years of European colonization, the generic heritage of the Garifuna people is down to only 20% Native American, yet their language is recognizably based on Awawak and Carib, with extensive borrowing from European languages as well as some West African influence.

Your claim that the ancient Europeans spoke an Amerind language is quite interesting. It is interesting because although we can reconstruct proto-languages I don’t know of any linguist who claims they can absolutely describe the languages spoken during the Neolithic. What is the linguistic evidence of a pan-Indian language spoken from Siberia to the Atlantic.

To the best of my knowledge there is no Amerind language related to Siberian. Please tell us the evidence supporting your findings.

One can't really rule out R1b's expansion with Megalithic cultures ca. 6000 BP to 4800 BP, but these cultures as so deeply intertwined with the first wave Neolithic in Europe, which was followed in many areas by population collapse (sometimes with reversion of foraging) after which there would be a window for resettlement by genetically distinct populations, that this seems less plausible to me.

Moreover, we also have two y-DNA samples from the Megalithic SOM-culture in France, dating to 2750-2725 BC. Both were I-M26.

Regarding the question whether the early R1b-M269 expansion occurred in a Basque-related population, that's an open question. But there appears to be some correlation between R1b-M269 and the autosomal Ancient North Eurasian admixture in Europe. It's just my impression, I didn't verify this correlation mathematically. But it would make sense, considering that Mal'ta boy was R*. If that correlation exists, we cannot treat the Basques as the epitomes of R1b-M269 people, because the Scottish, who have similarly high frequencies of R1b, are much more Ancient North Eurasian admixed than the Basques.

And let me just add one more thing: I agree that the Bell Beaker complex, at least in its majority, presumably wasn't IE speaking.But the assumption that R1b was widespread in the Bell Beaker culture stands on shaky grounds, especially since it wasn't anthropologically uniform. That was already evident from craniometry, and as Jocelyne Desideri has shown on the teeth, the people of the eastern group in the Czech republic and the Iberian group both had different local roots. (Although the evidence from mt-DNA certainly suggests some Iberian influence in central Europe.)

Dr. Clyde Winters wrote: "Your claim that the ancient Europeans spoke an Amerind language is quite interesting. It is interesting because although we can reconstruct proto-languages I don’t know of any linguist who claims they can absolutely describe the languages spoken during the Neolithic. What is the linguistic evidence of a pan-Indian language spoken from Siberia to the Atlantic.

To the best of my knowledge there is no Amerind language related to Siberian. Please tell us the evidence supporting your findings."

You're asking for quite a data dump!

Recently, linguists have accepted the link between Yeniseian languages and Dene languages of North America. I've already noted that Basque incorporates a lot of pronouns that appear to be derived from Dene languages. What's more, there also appears to be some Iroquoian influence in Basque as well. For instance, their words for 'he' (third person singular), jura, bera appear to be compound words incorporating the Iroquoian re- in addition to Dene pronouns. As another example, the Basque word for 'mouth', aho, closely resembles its Cherokee equivalent, aholi.

I've posted elsewhere in these blogs about how both personal and relative pronouns in IE and Iroquoian appear to be related - for instance, the word particles -an- / -on- denote third person plural pronouns in both Iroquoian languages and eastern IE languages (Indo-Iranian, Slavic).

Likewise, the words for 'water' in most IE languages appear to be derived from a common root from the word for word for 'water' in most Iroquouian languages. awa (Proto Indo-European 'PIE' akwa). (Some IE philologist have hypothesized that there were other root words for 'water' in IE but I they did not know about Yeniseian, and apparently mistook words derived from Kett ul and Kott ur as PIE.)

I am working my way down the Swadesh list, but I have found some interesting likely cognates down in the higher numbers between modern Iroouoian and PIE:

"Many researchers have committed on the large number of R-V88 who speak Afro-Asiatic languages. But few have noticed that many Niger-Congo speakers, Pgymies and Khoisan carry R-M269.

. Berniell-Lee et al (2009) found in their study that 5.2% carried Rb1*. The frequency of among the Bantu ranged from 2-20. The bearers of R1b1 among the Pygmy populations ranged from 1-25% (Berniell-Lee et al, 2009). The frequency of R1b1 among Guinea-Bissau populations was 12% (Carvalho et al,2010).Gonzalez et al, in The genetic landscape of Equatorial Guinea and the origin and migration routes of the Y chromosome haplogroup R-V88, found that 53& of the subjects in his study carried R-M269.

Given the Corded Ware/Beaker culture sites in North Africa, and widespread evidence of R-M269 from Guinea to South Africa offer the possibility that Sub-Saharan Africans may have introduced this haplogroup to western Eurasia."

I'm sorry but this is nonsense. The only place in Africa where R1b-M269+ and R1b-M269+ makes appears at very small frequencies is in North Africa, which was likely introduced there by European colonists. All those studies you mention about R1b*, which likely would mean R1b(xP25*) or R1b-M343 or R1b-M173, are pre-Cruciani.et.al.2010, and likely pre-R1b-V88 discovery. So just because they weren't typed for the V88 SNP they appeared ancestral. After Cruciani.et.al.2010 it became clear that all SubSaharan Africans are R1b-V88+ derived. In fact according to Cruciani.et.al.2010 they were only 5 people in their database with paragroup R1b-P25*, three Europeans, one West Asian, one East Asian if I recall correctly. So please, stop making absurd claims such as saying that haplogroup R originated in Africa. Or that West Africans lived in the Basque Country. Basques mt-DNA which has been well studied in ancient and modern samples lend no support to such theory, perhaps the discovery of mt-DNA L2 in Tres Montes Bronze Age Navarra could support some minor African influence, however the vast majority of Basque mt-DNA belongs to either mt-DNA H or U, with K, X, T making smaller counts.

This paper provides these statistics for R-M269 in Table 2, of the supplement. Anyone interested may check out the paper and supplement for themselves.Obviously you have not kept up with the literature on R-M269 in Africa.

Your comment about nonsense is clearly based on your lack of knowledge on the literature relating to R-M269 in Africa.

It is ludicris to believe that Europeans spread R-M269 into Africa, because whereas only 3% of the Mozabite Berbers and 28% of the Siwa carry this haplogroup, 53% of of the carriers of the R haplogroup in Guinea carry R-M269, this suggest that more people in Sub Saharan Africa carry this haplogroup than North Africans.

This makes my hypothesis that:"Given the Corded Ware/Beaker culture sites in North Africa, and widespread evidence of R-M269 from Guinea to South Africa offer the possibility that Sub-Saharan Africans may have introduced this haplogroup to western Eurasia;" highly possible, given the linguistic relationship between Basque and Niger-Congo language (Dogon).

African haplogroups have not only been found at Tres Montes Bronze Age Navarra, they were found in many ancient Iberian skeletons.

Seventy percent of the mtDNA of Sub-Saharan Africa belong to L,L2 and L3. Haplogroup L2 and L1b, is concentrated in western-central Africa, particularly along the coastal areas. Dominguez (2005) ,noted that much of the ancient mtDNA found in Iberia has no relationship to the people presently living in Iberia . Dominguez found that the lineages recovered from ancient Iberian skeletons are the African lineages L1b,L2 and L3. Almost 50% of the lineages from the Abauntz Chalcolithic deposits and Tres Montes, in Navarre are the Sub-Saharan lineages L1b,L2 and L3. The appearance of phylogenetically related sequences of hg L3 present in many ancient Iberian skeletons suggest that this haplogroup may have a long history in Iberia. This would support the possibility that West Africans were in ancient Iberia.

What I find rather striking is the strong predominance of y-haplogroup I2 in the late Bronze Age Lichtenstein cave in Germany. That sample is from the Urnfield culture around 1000 BC, and from central Europe. It's the only Bronze Age y-DNA sample we've got from central Europe. And it isn't dominated by R1b. I don't know if it's just atypical by accident, but that wouldn't be a reasonable assumption. So maybe there were really stronger pockets of R1b in the Western European Bronze Age than in central Europe. Maybe it really diffused in the Bell Beaker culture. It's not known how it got to western Europe. Maybe it was from the Yamnaya expansions, or from western Asia. But no matter where it came from, it ended up as the predominant haplogroup in non-IE Basques, who were presumably related with the Bell Beaker culture. If that is so, then it would follow that the ANE admixture in western Europe isn't directly linked with the expansion of R1b. Presumably R1b people were strongly ANE admixed in the beginning, but if my above reasoning is correct, they must have lost most of it by the time they reached western Europe. It's conceivable that the spread of R1b and of the ANE admixture were not closely linked. After all, Scandinavians are also rather strongly ANE admixed, yet they are dominated by haplogroup I1. So maybe the ANE admixture reached western Europe with the Celts, and from central Europe, but after R1b.

As for Grigoriev's theory to derive the Italo-Celts and the Germano-Balto-Slavs from West Asia via Iran, central Asia and Siberia: It occured to me that such a derivation would explain the Gedrosia component in the Italo-Celts and the Germanics. The Gedrosia component is particularly strong in northwestern Europe. But it's close to zero in Slavic and Baltic populations. These have got the Caucasus component instead. So therefore it's dubious that they could be derived in the same way. It makes more sense to derive them from the Corded Ware, because its mt-DNA showed links to the Caucasus area in the recent Brandt and Haak paper.

Well, the Basques still do have some ANE admixture according to Lazaridis et al., so it didn't get completely lost till R1b reached western Europe. And they are not shifted towards west Asia, which would speak against a West Asian origin of their R1b.

R1b indo European is going to be an elite takeover. The language is the language of the powerful, and trade. Similar to Latin, Greek, Aramaic, etc. same as how the Normans changes English and many other historical examples. The Italo Celto Germanic branch is the third oldest indo European branch. Easily old enough to spread with even bell beaker elite r1b.

Finding i2 in a Bronze Age site is expected. R1b would still be small. Even graves dating to 500ad in places have r1b second to Neolithic lineages. A 1-2% replacement per generation can explain today's numbers. It would only take a few dozen men 1500 years to nearly replace local lineages. Look at Niall of nine hostages. 1700 years and 21% of the region and 8% of all Irish possess this subclade.

Wild replacent theories are rediculous. We don't have a couple million victims of violent death, sorry!

I think you haven't quite understood my point. Actually I was doubting that R1b in western Europe has anything to do with Indo-Europeans, left alone an IE elite. Maybe R1b did spread from the Yamnaya culture into Europe. But the funny thing is that in central Europe, these R1b people apparently got assimilated by the Bell Beaker culture which originated in Iberia and presumably wasn't IE, at least not originally and predominantly. It's possible that this opened up the door for R1b in the non-IE Basques. Yes, R1b is predominant in western Europe, but this doesn't make it Italo-Celtic at its root. We have to be open to the possibility that it is the legacy of a non-IE substrate. That sometimes the substrate prevails goes without saying, see for instance Scandinavia and I1. I have said before that in southwestern Europe, R1b is actually more common in the areas where IE was introduced late, by the Romans (or not at all in the case of the Basques) than in the old Celtic and „para-Celtic“ areas. I think it's quite possible that the Yamnaya culture played a key role in the Indoeuropeanization of the Balkans, but it remains dubious that they were the Proto-IEs or the source of the Italo-Celts. I think most experts agree that Italo-Celtic split from the IE mainstream early, but this does in no way entail that it was also one of the earliest IE arrivals in central Europe. That would be a logical fallacy. According to Stanislav Grigoriev's theory it arrived late, with the Tumulus culture, around 1600 BC, from Siberia. It has been observed before that Italo-Celtic displays similarities with Tocharian. From this, some have concluded that Tocharian originated in central Europe. Others have suggested that the similarities are merely the consequence of the early spin-off of both branches. But Gamkrelidse and Ivanov suggested that there really once was a Tocharo-Celto-Italic branch, besides Anatolian and the still undifferentiated rest. In my view such an eastern origin of Italo-Celtic would explain the pattern of the Dodecad K12b Gedrosia admixture in Europe. The Gedrosia component is the eastern part of the more general West Asian component, the western part being the Caucasus component. And as I said, with the middle bronze age we have the first firm evidence for horse riding and chariots in Europe. The origin of the Tumulus culture isn't quite clear afaik. It has been suggested that it is the result of a push from the Danubian area. Later archaeologists preferred to look for local continuities, under the general trend of antimigrationism. The Polish wikipedia article claims that its origins are sought more in the western parts of its distribution area. But to my knowledge it's slightly older in the eastern parts, in Bohemia, Moravia and Lower Austria. And apparently its introduction and spread involved violence.

Finding I2 completely predominating in a sample from late Bronze Age central Europe, from the Urnfield culture, isn't quite expected, at least under the paradigm that the Italo-Celts were originally R1b. Few people will dispute that the Urnfield culture was a major source and early florescence of Italo-Celtic tribes.

As for your calculation of the slow, gradual implementation of R1b with a steady, low replacement rate, the question is, why should one haplogroup be advantaged across 15 centuries? It's dubious that social forces could act into the same direction for such a long time. A selective advantage would be needed...

Even graves dating to 500ad in places have r1b second to Neolithic lineages.

Ha, yes for sure, but not in central – western Europe! A Basque sample from that time was already largely R1b. I wonder what sample you are referring to??

Lastly, while I didn't propagate any wild replacement theories, it's clear from the facts (Lazaridis et al. 2013), that there was a non-negligible admixture from an Ancient North Eurasian admixed population in post-mesolithic and post-early neolithic times. It's of course possible that British hunter-gatherers were already more ANE-admixed than those in Scandinavia and Luxemburg. But this isn't a plausible scenario at all.

This paper provides these statistics for R-M269 in Table 2, of the supplement. Anyone interested may check out the paper and supplement for themselves.Obviously you have not kept up with the literature on R-M269 in Africa.

Your comment about nonsense is clearly based on your lack of knowledge on the literature relating to R-M269 in Africa.

It is ludicris to believe that Europeans spread R-M269 into Africa, because whereas only 3% of the Mozabite Berbers and 28% of the Siwa carry this haplogroup, 53% of of the carriers of the R haplogroup in Guinea carry R-M269, this suggest that more people in Sub Saharan Africa carry this haplogroup than North Africans.

Equatiorial Guinea was a colony of Spain, and by all means those R-M269 could be descendants of Spaniards. Again, with the exception of North Africa, there is no R-M269 derived haplogroups in Sub-Saharan Africa. Moreover, there isn’t any R1b-M343 in Sub-Saharan Africa, nor is there any R1a derived groups in Sub-Saharan Africa. Take a look at what the authors of the study say relating to R1b-M269 in Equatorial Guinea: Altogether, the proportion of recent Eurasian admixture found in our sample is approximately 15% (haplogroups E1b1b1b-M81, G-M201, N1c-Tat, R1b1b2-M269 and two chromosomes belonging to E1b1b1a-M78—see criteria below), which is easily explained by the well-reported European arrivals to this territory within the last five centuries.So your argument that whenever a Eurasian haplogroup is found in Sub-Saharan Africa it denotes a potential origin in the region is not only fallacious, but completely defies logic, and Occam’s Razor. Do you know how many hoops you have to jump to really make R1b-M269 be born in Africa. What it shows in fact to me, and other readers experienced with the phylogenetics of R1b-M269 is that you lack complete and total understanding of the genetic structure of such haplogroup. What is worse is that you claim that haplogroup R* the parent of R1b and R1a was born in Africa. Again some of your claims are you downright absurd and fall in the realm of trolling. Easy proof for a non African origin of R1b-M269:

1-Older observed genetic diversity in Europe and the Middle East(Balaresque.et.al.2010, Myres.et.al.2011, Cruciani.et.al.2010). Lack of parental subclades in Africa. R1b-P25, R1b-M343. Autosomal DNA does not suggest a recent(<5000 years) signature of African admixture in populations that are heavy in R1b-M269 such as the Basques, Irish, English, French, etc. Ancient DNA shows both R1b and R1a in Europe 5000 years ago, it shows R in Eurasia 20,000 years ago. So once more, I will kindly request you stop spreading such nonsense on the web.

.“Seventy percent of the mtDNA of Sub-Saharan Africa belong to L,L2 and L3. Haplogroup L2 and L1b, is concentrated in western-central Africa, particularly along the coastal areas. Dominguez (2005) ,noted that much of the ancient mtDNA found in Iberia has no relationship to the people presently living in Iberia . Dominguez found that the lineages recovered from ancient Iberian skeletons are the African lineages L1b,L2 and L3. Almost 50% of the lineages from the Abauntz Chalcolithic deposits and Tres Montes, in Navarre are the Sub-Saharan lineages L1b,L2 and L3. The appearance of phylogenetically related sequences of hg L3 present in many ancient Iberian skeletons suggest that this haplogroup may have a long history in Iberia. This would support the possibility that West Africans were in ancient Iberia.”

Not a single one of them found mt-DNA L in ancient Basque samples(n=49). In fact you are using an outdated thesis who only made use of HVR-I to identify haplogroups, and also, it is 50% of 4, meaning the sample size was extremely small, so 2 out of 4 while being 50% doesn’t change the fact that 49 other ancient samples haven’t reproduce the same results. Also here is another study, using different sites featuring a total of 116 ancient dna samples:

You claim that R-M269 in Guinea is due to mating with Spanish people, this is pure speculation. This is a false theory , like the view that all African haplogroups in Southern Europe are the result of the slave trade.Have you forgotten that West Africa, was considered the "European" graveyard.

Secondly, I did not say the Basque carried L haplogroups, I said there was evidence of Iberians carring L haplogroups in ancient times. Stop trying to flip the script to make yourself correct.

The widespread existence of R-M269 can not be explained by the mixing of Africans and Europeans.

By the way, judging from the Eupedia maps, y-haplogroup I2b, which was once believed to peak in northern Germany, is quite frequent (3 - 4%) in northwestern France and Cornwall, and even more common (5 – 6%) in southwestern Scotland and the Scottish Lowlands, and in large parts of Ireland (except for the southwest). This is probably a trace of the Celts who brought this haplogroup from their central European staging point.

A sceptic might say, I2 being very old, it may be anything, there's no reason to associate it with Celtic influence. However I note that, in NW-Europe, it doesn't correlate exclusively with Germanic influence but is also present at similar levels where Germanic influence was minimal. And moreover it doesn't correlate with R1b, to the contrary. While I2b is strong in the Scottish Lowlands, R1b is stronger in the Highlands. While I2b is strong in all of Ireland except for the southwest, R1b gets a bit weaker towards the northeast and east. These distribution patterns suggest that I2b arrived after R1b; R1b is particularly strong in refuge areas. At one point R1b must have arrived from the east, too, via central Europe presumably, but its enormous expansion appears to have occured after its arrival in western Europe. In other words: There was no R1b-population invading western Europe, it seems to have seeped in unspectacularly and expanded by chance because of a few successful males.

On a second thought, according to linguists there were several waves of Celtic into NW Europe, so it may be simply the case that a later wave had more I2b than previous ones. I think that the Gedrosia admixture in NW Europe has to be explained with an eastern, south Siberian origin of the Italo-Celts, in line with Grigoriev's theory. The problem is just: The European peak of Gedrosia lies in Argyll, and judging from the MDLP World-22 results, the Welsh are at least equally Gedrosia admixed. Yet the Welsh have close to 0% R1a, so how could they be admixed with an ancient South Siberian element? The solution can only lie in R1b.The basic branches of R1b are:V88 = Levant, AfricaM335 = AnatoliaP297 = ? It split into:M73 = Hazaras, Bashkirs, i.e. an eastern branchand into M269* = Balkans, Armenia, i.e. a western branchInterestingly M269* isn't purely western, but is also present in 2.4% of Bashkirs. So it must have been present in the West Asian ancestors of the Bashkirs, too. From M269* descended L23*, whose distribution is similar to that of M269*, just a little wider. It isn't only common in the Valais in Switzerland, but also in Kosovo, Armenia and in Bashkirs! So it too must have existed at the time when the West Asian ancestors of the Bashkirs started to migrate eastwards.So this L23* had a wide belt-shaped distribution, ranging from C Europe, across the Balkans, West Asia and from there through central Asia to the Urals, where the Bashkirs live.Gamkrelidze and Ivanov suggested that there once was a common Tocharo-Celto-Italic, besides Anatolian and the still undifferentiated rest.And there is increasing evidence that the forebears of the Tocharians arrived there from western Asia via the Silk Road, rather than being an eastern extension of Pontic-Caspian Kurgan tribes. Although the Tarim mummies were R1a, R1b is present in the Uighurs who live in the area now. The Tocharian language may have been brought by R1b people from western Asia. And Tocharian has a rich agricultural vocabulary which doesn't fit with a steppe origin.To me it's plausible that the Tocharo-Celto-Italic branch migrated from western Asia along the Silk Road and was rich in R1b-L23* and the Gedrosia component. Grigoriev has suggested that the forebears of the Italo-Celts, which he sees in the Tumulus culture have originated between the Urals and the Irtysh. From there they may have brought R1b-L23* and the Gedrosia admixture. It should also be noted that recent evidence has indicated that western European hunter-gatherers, early European farmers and Kurgan people from the Pontic steppe all were dark pigmented (except for light eyes in western hunter-gatherers). Genetic evidence of Bronze Age South Siberians however showed that they were light-pigmented. So this would also explain the light-mixed pigmentation of Celtic populations. And these immigrants would have boosted the North European component in NW and C Europe. I also point to mt-DNA haplogroup K2b. This was found in Bronze Age south Siberians. It is now common in C, N and W Europe. But there the question arises what about the Bell Beaker people of Kromsdorf? They were already R1b at a much earlier date! Well, that's correct, but as I had pointed out, R1b-L23* had a wide, belt-shaped distribution that included C Europe and the Balkans. So it's no surprise that it was found in German Bell Beaker people. But there is no evidence that these already spoke Italo-Celtic. Also, there is no evidence that M412 and the subsequent haplogroups descended from them. And what about the negative correlation of R1b with IE languages in SW Europe? I've once read that isolated, small populations are more susceptible to drift, and that mutations can get more easily fixed there. So R1b may have drifted more in the isolated Iberians and Basques.

Genetiker has alerted me of two things: R1b-M269* is also present in 5.3% of Germans, so it had already reached central Europe, too. And, more importantly: Current age estimates of R1b-L11 place it at 6.1 kya. This would strongly contradict my theory that it is descended from south Siberians who arrived in central Europe during the middle Bronze Age. There is also another, a linguistic objection: The diversity of Italo-Celtic languages makes it hard to believe that they reached central Europe as late as 1600 BC. There were two quite different branches in Italy, a p-Italic (Oscan-Umbrian) and a q-Italic (Latino-Faliscan), similar to the disctinction of p- and q-branches in Celtic. Plus there were related languages that diverged even more, like Ligurian and Lusitanian. So that theory of mine isn't really tenable. However, recently I noted that the difference between the Gedrosia admixture and the Caucasus admixture in Europe is mainly a slight Kalash relatedness of the Gedrosia component. And this explains its similarity with the MDLP World-22 Indo-Iranian component, which is mainly a Kalash component. Now, it occured to me that this influence seems correlated with R1b-L23. In the Caucasus the World-22 Indo-Iranian component is strongest in Tabassarans (3.8%), followed by Lezgins (2.9%). The Tabassarans have 37.2% R1b-L23*, and the Lezgins have 12.9%. The Indo-Iranian component is also relatively strong in Bashkirs (2%), and they have 32.2% L23*. In Europe, the Indo-Iranian component is strongest in Norwegians (3.8%), Orcadians (3.7%) and the Welsh (3.4%). They all have lots of L23, especially the Welsh, though mostly more derived. Notably the Indo-Iranian component is also present in the Basques (1.1%) which parallels the Gedrosia presence there. And BTW, this all parallels and explains the mysterious Dagestan component of the early Dodecad experiments. So, I would suggest that both R1b-L23* and some slight Kalash-like admixture entered southeastern Europe early, from West Asia / eastern Anatolia, presumably with the chalcolithic Baden culture. Presumably this didn't carry yet a lot of general West Asian admixture. It has been suggested that the domestic pottery of the eastern Bell Beaker group is derived from Vucedol pottery, and this is derived from Baden. So it's all connected: R1b, Gedrosia, Bell Beaker pottery and most probably also the Dinaric Bell Beaker skulls. And, most likely Celtic languages. Bell Beaker-like culture and people continued to dominate southern Germany in the early Bronze Age. In the Middle Bronze age they were pushed to the west, by the invasive Tumulus culture which brought more Germanic-like languages.

It should be noted that, for several hundred years, the Berber corsairs raided heavily for slaves along the western coasts of Ireland and Scotland, with several raids traveling as far as Iceland and Norway. Said slaves might well have had fairly high status,as North Africans place a major emphasis on fair complexion.

Also, R1b/M269 males have a much higher sperm count and sperm motility than other groups, presumably due to selection resulting from a near constant state of war for millenia. If a group of men only have a few years to sow their seed, selection tends to favor the most fertile. This could well create an historical drift toward M269.

It all seems straightforward to me.There is no relation between SNP-hts and STRs, they are independent. For different hts to have different STRs, the descendants must have been very heavily pruned over an extended period so that only a few lines remain from ancient times.

This pruning has not occurred in Europe since the Bronze age - the population has grown steadily. Therefore no hts that have arisen in that time can be distinguished by STRs.

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